Research Article |
Corresponding author: Stuart V. Nielsen ( stunie@gmail.com ) Academic editor: Angelica Crottini
© 2020 Stuart V. Nielsen, Werner Conradie, Luis M. P. Ceríaco, Aaron M. Bauer, Matthew P. Heinicke, Edward L. Stanley, David C. Blackburn.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Nielsen SV, Conradie W, Ceríaco LMP, Bauer AM, Heinicke MP, Stanley EL, Blackburn DC (2020) A new species of Rain Frog (Brevicipitidae, Breviceps) endemic to Angola. ZooKeys 979: 133-160. https://doi.org/10.3897/zookeys.979.56863
|
Recent molecular phylogenetic work has found that Breviceps Merrem, 1820 comprises two major clades, one of which, the B. mossambicus group, is widely distributed across southern sub-Saharan Africa. This group is notable for harboring abundant cryptic diversity. Of the four most recently described Breviceps species, three are members of this group, and at least five additional lineages await formal description. Although Breviceps has long been known to occur in Angola, no contemporary material has been collected until recently. The three most widespread taxa, B. adspersus, B. mossambicus, and B. poweri, may all occur in Angola, but accurate species assignment remains challenging given the rampant morphological similarity between these taxa, and, until recently, the lack of genetic resources. Phylogenetic, morphological, and acoustic analyses of recently collected samples from disparate localities within Angola provide evidence for an undescribed species that is sister to B. poweri. The new species can be diagnosed from its sister taxon by lacking pale spots along the flanks, a pale patch above the vent, and a short, dark band below the nares (all present in B. poweri). Additionally, the male advertisement call differs from the three other Breviceps that might occur in Angola in having both a longer interval between consecutive calls and a higher average dominant frequency. We here describe this lineage as a distinct species, currently only known from Angola, and discuss the presence of other Breviceps taxa within Angola.
Investigações moleculares recentes revelaram que o género Breviceps Merrem, 1820, é composto por duas linhagens principais, uma das quais, o grupo B. mossambicus, é amplamente distribuído na região sul da África subsaariana. Este grupo é notável por albergar uma abundante diversidade críptica. Das quatro espécies de Breviceps recentemente descritas, três pertencem a este grupo, e pelo menos outras cinco linhagens adicionais aguardam a sua descrição formal. Apesar de o género ser conhecido de Angola desde há muito tempo, só muito recentemente foram colhidos novos espécimes. Os três taxa mais amplamente distribuídos, B. adspersus, B. mossambicus e B. poweri podem todos, porventura, ocorrer em Angola, no entanto a correta identificação destas espécies têm sido problemática devido às semelhanças morfológicas extremas entre este taxa, e, até muito recente, a completa ausência de material genético. Análises filogenéticas, morfológicas e acústicas dos espécimes recentemente colhidos em diferentes locais de Angola apontam para a existência de uma espécie nova para a ciência, irmã de B. poweri. A nova espécie pode ser diferenciada do seu táxon irmão pela falta de marcas pálidas nos flancos, mancha pálida acima do ventre e pequena banda negra abaixo do nariz (presentes em B. poweri). Para além destas características, o chamamento dos machos difere das outras três espécies de Breviceps que podem ocorrer em Angola por ter um maior intervalo entre chamamentos consecutivos e uma maior frequência média dominante. Descrevemos aqui esta linhagem como uma espécie distinta, atualmente apenas conhecida de Angola, e discutimos a presença de outras espécies de Breviceps em Angola.
Afrobatrachia, Anura, Breviceps ombelanonga sp. nov., cryptic species, multilocus, novel species, Sub-Saharan Africa
África Subsahariana, Afrobatrachia, Anura, Breviceps ombelanonga sp. nov., espécies crípticas, espécies novas, multilocus
Breviceps Merrem, 1820 is a genus of fossorial frogs widely distributed across southern sub-Saharan Africa, from Angola in the northwest, through Zambia, the southern portions of the Democratic Republic of the Congo and Tanzania, and southward throughout virtually all of southern Africa (Minter 2004;
The taxonomy of Angolan Breviceps has long been problematic.
A recent phylogenetic study of Breviceps (
Angola’s long civil war, which lasted from 1975 to 2002, effectively stifled biological exploration and discovery (for additional summary, see
We consider species as units of separately evolving metapopulation lineages, following the conceptual framework developed by
Between 2016 and 2019, specimens referable to the genus Breviceps were collected from three main localities within Angola (Fig.
Sampling information including specimens/field IDs (Museum Abbreviations:
species | Tree ID | Specimen ID | Field ID | Latitude and Longitude | Country | Locality | RAG1 | BDNF | 12S | 16S |
---|---|---|---|---|---|---|---|---|---|---|
Breviceps ombelanonga sp. nov. | ANG-01 | UF Herp 187172 |
|
-9.183833, 13.371472 | ANG | Kawa Camp (1 km S of the Kwanza R.), Kissama NP, Luanda Prov. | MT944215 | MT944224 | MT944230 | MT944241 |
ANG-02 | UF Herp 187173 |
|
-9.183833, 13.371472 | ANG | Kawa Camp (1 km S of the Kwanza R.), Kissama NP, Luanda Prov. | MT944216 | MT944225 | MT944231 | MT944242 | |
ANG-03 | MHNCUP_ANF 0320 | AMB11736 | -11.083845, 16.667410 | ANG | Embala Seque (14 km N of Cassumbi village), Bie Province | MT944217 | MT944226 | MT944232 | MT944243 | |
ANG-04 |
|
WC-3924 | -12.689351, 18.360115 | ANG | Cuito River source lake, Moxico Province | MT944218 | MT944227 | MT944233 | MT944244 | |
ANG-05 |
|
WC-4591 | -13.089343, 18.894850 | ANG | Cuanavale River source lake, Moxico Province | MT944219 | MT944228 | MT944234 | MT944245 | |
ANG-06 |
|
WC-4756 | -13.135440, 19.043970 | ANG | Quembo River source lake, Moxico Province | MT944220 | MT944229 | MT944235 | MT944246 | |
ANG-07 |
|
WC-4827 | -13.003340, 19.135640 | ANG | Cuando River source, Moxico Province | – | – | MT944236 | MT944247 | |
B. adspersus | ADS-01 |
|
AMB8318 | -22.708056, 29.528333 | RSA | Farm Celine, Limpopo | MT944221 | – | MT944237 | MT944248 |
ADS-02 |
|
|
-18.670972, 26.953472 | ZIM | Hwange | MT944222 | – | MT944238 | MT944249 | |
ADS-03 |
|
|
18.628793, 26.872087 | ZIM | Miombo Safari Camp | MT944223 | – | MT944239 | MT944250 | |
ADS-04 |
|
|
-19.528500, 17.564167 | NAM | Farm Ohange, Otjozondjupa | – | – | MT944240 | MT944251 | |
ADS-05 | – | SVN 766 | -23.731926, 27.579803 | RSA | Ellisras | MH340062 | MH340138 | MH340291 | MH340369 | |
ADS-06 | – | SVN 768 | -23.731926, 27.579803 | RSA | Ellisras | MH340063 | MH340139 | MH340292 | MH340370 | |
ADS-07 |
|
AMB7963 | -17.623556, 24.199583 | NAM | Katima Mulilo | MH340064 | MH340140 | MH340293 | MH340371 | |
ADS-08 | – | AMB7972 | -18.000000, 21.070000 | NAM | Caprivi | MH340065 | MH340141 | MH340294 | MH340372 | |
ADS-09 |
|
AMB7980 | -18.035500, 20.971528 | NAM | Caprivi | MH340066 | MH340142 | MH340295 | MH340373 | |
B. mossambicus | MOS-01 |
|
DMP 344 | -15.463942, 36.977847 | MOZ | Gurue | MH340075 | MH340151 | MH340304 | MH340382 |
MOS-02 |
|
|
-15.933333, 35.516667 | MW | Mulanje | MH340076 | MH340152 | MH340305 | MH340383 | |
MOS-03 |
|
RB09-159 | -15.030944, 40.740944 | MOZ | Ila de Mozambique | MH340077 | MH340153 | MH340306 | MH340384 | |
MOS-04 |
|
RB09-179 | -15.030944, 40.740944 | MOZ | Ila de Mozambique | MH340078 | MH340154 | MH340307 | MH340385 | |
MOS-05 |
|
RB09-030 | -12.963611, 40.529444 | MOZ | Pemba | MH340079 | MH340155 | MH340308 | MH340386 | |
MOS-06 |
|
RB09-046 | -12.963611, 40.529444 | MOZ | Pemba | MH340080 | MH340156 | MH340309 | MH340387 | |
MOS-07 |
|
RB10-A097 | -15.030722, 40.741222 | MOZ | Nampula | MH340081 | MH340157 | MH340310 | MH340388 | |
MOS-08 |
|
NIMB 112 | -13.308060, 35.244114 | MOZ | Lichinga | MH340082 | MH340158 | MH340311 | MH340389 | |
MOS-09 |
|
Syran 12 | -13.288667, 38.681528 | MOZ | Balama | MH340083 | MH340159 | MH340312 | MH340390 | |
B. poweri | POW-01 | – | ELI 325 | -7.277700, 27.389800 | DRC | Manono | MH340084 | MH340160 | MH340313 | MH340391 |
POW-02 |
|
JWH10-A114 | -12.237778, 25.341944 | ZAM | Kalumbila | MH340085 | MH340161 | MH340314 | MH340392 | |
POW-03 |
|
RB10-F003 | -15.510278, 28.260528 | ZAM | Lusaka | MH340086 | MH340162 | MH340315 | MH340393 | |
POW-04 |
|
RB10-F012 | -15.510278, 28.260528 | ZAM | Lusaka | MH340087 | MH340163 | MH340316 | MH340394 |
We amplified partial sequences of two mitochondrial (12S and 16S ribosomal rRNA genes) and two nuclear loci (recombination activating protein 1, RAG1; brain derived neurotrophic factor, BDNF) using the PCR primers and cycling conditions outlined in
Uncorrected mean pairwise 12S and 16S mitochondrial sequence differences between ingroup Breviceps sequence pairs (above/below the diagonal, respectively) and within species (along the diagonal) conducted in MEGA.
16S/12S | 12S | ||||
B. ombelanonga sp. nov. | B. adspersus | B. poweri | B. mossambicus | ||
16S | B. ombelanonga sp. nov. | 0.04/0.03 | 0.09 | 0.09 | 0.11 |
B. adspersus | 0.11 | 0.01/0.01 | 0.09 | 0.08 | |
B. poweri | 0.12 | 0.11 | 0.02/0.02 | 0.09 | |
B. mossambicus | 0.12 | 0.08 | 0.10 | 0.01/0.01 |
Datasets (concatenated and partitioned by locus/codon) of all samples were analyzed using maximum likelihood (RAxML v.8.2;
Specimens were measured to the nearest 0.1 mm using digital calipers under a dissecting stereomicroscope for the following 24 morphological characters as defined by
Advertisement calls were recorded in the field using an Samsung Galaxy Note 3 cellphone at a sampling rate of 44100 kHz, and analyzed using Sound Ruler Acoustic Analysis v.0.9.6.0 using default settings (
The electronic edition of this article conforms to the requirements of the amended International Code of Zoological Nomenclature (ICZN), and hence the new names contained herein are available under that Code from the electronic edition of this article. This published work and the nomenclatural acts it contains have been registered in ZooBank, the online registration system for the ICZN. The ZooBank LSIDs (Life Science Identifiers) can be resolved and the associated information viewed through any standard web browser by appending the LSID to the prefix “http://zoobank.org/”. The LSID for this publication is: http://zoobank.org/References/2043280A-1591-4D51-ACE3-F9015F170890. The electronic edition of this work was published in a journal with an ISSN, and has been archived and is available from the following digital repositories: PubMed Central, LOCKSS.
Our concatenated, multi-locus dataset was 1,852 bp long, of which 390 characters were parsimony informative. Phylogenetic analyses resulted in a well-supported species-level phylogeny and high support that Breviceps is monophyletic (bootstrap [bs] 100, posterior probability [pp] 1.0; Fig.
Geographic distribution and phylogenetic relationships of Breviceps spp. included in this study. A Map of Angola and surrounding countries with all known Breviceps spp. sampling localities indicated on legend. The proposed distributions of B. adspersus and B. poweri (blue and red polygons, respectively) are from
Mensural and meristic data are presented in Table
PCA plots of ten size-corrected morphological characters from specimens of B. adspersus (n = 24), B. mossambicus (n = 9), B. poweri (n = 8), and the putative new Angolan species (n = 6) (Suppl. material
UF Herp 187172 | UF Herp 187173 | MHNCUP_ANF 0320 |
|
|
|
|
|
Average | SD | |
---|---|---|---|---|---|---|---|---|---|---|
SVL | 30.5 | 27.5 | 24.6 | 25.4 | 18.3 | 26.6 | 30.1 | 26.5 | 26.2 | 3.80 |
SUL | 26.3 | 24.9 | – | 23 | 17.6 | 23.2 | 29 | 25.2 | 24.2 | 3.53 |
HL | 7.2 | 7.3 | 7.3 | 10.8 | 6.4 | 9.6 | 12.7 | 9.3 | 8.8 | 2.17 |
ES | 3.2 | 2.8 | 2.9 | 3.1 | 2.2 | 2.7 | 3.3 | 2.5 | 2.8 | 0.37 |
NOD | 2.0 | 1.7 | 1.6 | 2.1 | 2.5 | 1.7 | 2.2 | 2.1 | 2.0 | 0.30 |
ED | 2.7 | 2.8 | 3.2 | 3.4 | 1.3 | 3.1 | 3.6 | 2.8 | 2.9 | 0.70 |
NLD | 1.7 | 1.7 | 1.3 | 1.4 | 1 | 1.2 | 1.8 | 1.3 | 1.4 | 0.28 |
ELD | 1.7 | 1.3 | 1.7 | 1.8 | 1.4 | 2.1 | 2.3 | 1.9 | 1.8 | 0.33 |
IND | 1.9 | 1.9 | 2 | 1.6 | 1.4 | 1.8 | 2.2 | 2 | 1.9 | 0.25 |
MW | 7.2 | 6.8 | 5.1 | 6.9 | 4.7 | 6.3 | 7.9 | 7 | 6.5 | 1.08 |
EAD | 9.1 | 9.3 | 4.1 | 4.5 | 3.6 | 4.3 | 5.3 | – | 5.7 | 2.42 |
F1L | 1.7 | 1.5 | 1.5 | 2.7 | 1.7 | 2.7 | 2.5 | 2.8 | 2.1 | 0.59 |
F2L | 1.9 | 1.9 | 1.7 | 3.6 | 1.8 | 3.2 | 2.6 | 2.7 | 2.4 | 0.71 |
F3L | 2.9 | 3.0 | 2.2 | 3.9 | 2.6 | 3.9 | 3.6 | 3.7 | 3.2 | 0.64 |
F4L | 1.2 | 1.3 | 1.2 | 1.8 | 0.9 | 2 | 1.5 | 1.4 | 1.4 | 0.35 |
T1L | 1.0 | 1.1 | 1 | 1.4 | 0.6 | 1.5 | 1.6 | 1.2 | 1.2 | 0.32 |
T3L | 1.8 | 1.9 | 2.9 | 2.4 | 2 | 2.7 | 1.5 | 2.7 | 2.2 | 0.51 |
T4L | 4.4 | 4.2 | 4.6 | 4.1 | 3.2 | 5 | 4.9 | 4.6 | 4.4 | 0.57 |
FT | 10.9 | 10.4 | 8 | 8.9 | 6 | 10.3 | 10.5 | 9.7 | 9.3 | 1.65 |
T5L | 4.0 | 5.0 | 0.8 | 4 | 3.2 | 4.9 | 5.3 | 4.7 | 4.0 | 1.46 |
MTL | 1.0 | 1.0 | 1.2 | 2.3 | 1.7 | 2.3 | – | – | 1.6 | 0.61 |
IMTL | 3.0 | 2.9 | 3 | 3.2 | 2.3 | 3.4 | 3.6 | 3.7 | 3.1 | 0.45 |
TIB | 8.5 | 8.3 | – | 6.4 | 4.8 | 7.4 | 8.8 | 8.1 | 7.5 | 1.43 |
The advertisement call of the eastern population is pulsed, has a call duration of 0.175 ± 0.083 s, with relatively long intervals between consecutive calls (0.996 ± 0.133 s), a high number of pulses per call (28–34; Table
Principal components analysis (PCA) loadings based on 10 size corrected morphological characters (head length, HL; head width, HW; eye diameter, ED; snout length, ES; interorbital distance, IOD; internarial distance, IND; thigh length, THL; crus length, CL; foot length, FL; and length of manual digit III, F3L).
PC1 | PC2 | PC3 | PC4 | SShapiro-Wilks test | |
Proportion of Variance | 29.92 | 26.57 | 13.70 | 8.67 | |
Cumulative Proportion | 29.92 | 56.49 | 70.19 | 78.86 | |
Loadings | |||||
Head Length (HL) | -0.3025245 | 0.35446298 | -0.3789378 | 0.02808779 | W = 0.943, p = 0.083 |
Head Width (HW) | 0.33281314 | 0.38971121 | 0.08705045 | 0.33211061 | W = 0.956, p = 0.205 |
Eye diameter (ED) | -0.2210569 | 0.22060777 | -0.6170786 | -0.1165699 | W = 0.963, p = 0.312 |
Snout length (ES) | -0.4530121 | -0.0316756 | 0.28394189 | -0.1665634 | W = 0.973, p = 0.567 |
Interorbital distance (IOD) | -0.3043034 | 0.24152256 | 0.52046607 | 0.21274977 | W = 0.959, p = 0.240 |
Internarial distance (IND) | -0.4018886 | 0.1543163 | 0.09759718 | 0.52889612 | W = 0.965, p = 0.360 |
Thigh length (THL) | 0.2280386 | 0.3347664 | 0.24257868 | -0.4343642 | W = 0.965, p = 0.360 |
Crus length (CL) | 0.45829394 | 0.12745601 | 0.05499745 | 0.20448849 | W = 0.975, p = 0.636 |
Pes length (FT) | -0.1152479 | 0.43230304 | 0.17838021 | -0.5130485 | W = 0.900, p = 0.005 |
Manual digit III length (F3L) | 0.11914774 | 0.52484652 | -0.115415 | 0.16994577 | W = 0.989, p = 0.978 |
Our phylogenetic analyses indicate that sampled individuals from Angola form a clade that is genealogically exclusive from other described species of Breviceps (Fig.
Breviceps gibbosus:
Breviceps gibbosus:
Breviceps mossambicus:
Rana mossambicus: Hellmich (1957: 30).
Breviceps “mossambicus-adspersus” complex:
Breviceps adspersus [part]:
Breviceps sp.:
Breviceps cf. adspersus:
Holotype. UF Herp 187172 (field number
A species referable to Breviceps due to the following characteristics (
The new species can be distinguished from other species of Breviceps occurring in the region by the following: pale paravertebral and dorsolateral patches are lacking, although a fine dorsolateral band may be present (versus no pale paravertebral or dorsolateral spots or patches in B. mossambicus; series of both paravertebral and dorsolateral pale spots and patches present in B. adspersus, a series of pale dorsolateral spots or patches present in B. poweri); no conspicuous light patch above vent (present in B. poweri); manual digit IV reaching approximately midway between the proximal and distal subarticular tubercles of manual digit III (versus nearly reaching distal subarticular tubercle of manual digit III in B. mossambicus; not reaching or barely passing the proximal subarticular tubercle of the manual digit III in B. poweri; similar to B. adspersus in usually not reaching the distal subarticular of manual digit III); gular region with a single uniformly dark patch (versus a pair of marbled to freckled patches in B. adspersus).
The advertisement call of the new species (Table
Comparison of the main variables for the advertisement calls of Breviceps ombelanonga sp. nov., Breviceps mossambicus, Breviceps adspersus and Breviceps poweri. Comparative data taken from
B. ombelanonga sp. nov. | B. adspersus | B. mossambicus | B. poweri | |||||
---|---|---|---|---|---|---|---|---|
avg ± sd | range | avg ± sd | range | avg ± sd | range | avg ± sd | range | |
Call duration (s) | 0.175 ± 0.083 | 0.064–0.342 | 0.196 ± 0.047 | 0.077–0.293 | 0.500 ± 0.070 | 0.036–0.079 | 0.140 ± 0.012 | 0.111–0.160 |
Call interval (s) | 0.996 ± 0.133 | 0.742–1.190 | 0.745 ± 0.636 | 0.363–0.745* | 0.710 ± 0.168 | 0.396–1.17 | 0.743 ± 0.166 | 0.500–1.100 |
No. of pulses/call | 30 ± 2.6 | 28–34 | 23 ± 3.3 | 14–31 | 9 ± 1.2 | 7–13 | 30 ± 16.3 | 10–74 |
Dominant frequency (Hz) | 2156 | na | 1742 ± 100 | 1482–2179 | 1835 ± 107 | 1600–2193 | 1728 ± 83 | 1557–1903 |
Adult male (SUL 30.5 mm), with globular body and well-developed short limbs with medialmost and lateralmost digits reduced (Fig.
Skin of dorsum and head smooth, and weakly glandular with irregular folds; skin of ventrum smooth; skin folds overlying vent creating triangular shape.
Limbs short with digits I and V short or rudimentary; webbing absent on manus and pes; nuptial pads absent and adhesive glands not discernable; relative manual digit lengths when adpressed: III>II>I>IV; only tip of first pedal digit extending beyond fleshy webbing and sole; fourth (outer) manual digit reaches midway between the large tubercle at metacarpophalangeal joint and subarticular tubercle at most proximal interphalangeal joint; finger tips conical, not expanded; several small globular palmar tubercles; single subarticular tubercles present on pedal digits II, III, and IV; pedal digit V very short, falling short of most proximal subarticular tubercle of pedal digit IV; well-developed (though not keratinized) inner metatarsal tubercle visibly longer than pedal digit III, separated from conical outer metatarsal tubercle by deep cleft.
In life, dorsum of body mottled dark brown on pale tan base, transitioning to golden yellow on the lateral aspects, before stark transition to solid dark brown flanks with a dark boundary becoming paler ventrally (Fig.
In preservative, coloration is largely similar but more muted and overall darker (Fig.
Measurements of the type series are shown in Table
All specimens resemble the holotype in the absence of a visible tympanum, and skin that is densely granular dorsally and laterally and smooth ventrally (Figs
Variation in color and pattern within living paratypes of B. ombelanonga sp. nov.: A, B sub-adult (of unknown sex) from Embala Seque (14 km N of Cassumbi village), Bié Province (MHNCUP_ANF 0320) C juvenile male, Cuito River source lake, Moxico Province (
Color and pattern in UF Herp 187173 is very similar to the holotype. Dorsum gray with scattered black spots (
Variation in color and pattern within preserved paratypes of B. ombelanonga sp. nov.: A an adult male from Kawa Camp Headquarters, Luanda Province (UF Herp 187173) B sub-adult (of unknown sex) from Embala Seque, Bié Province (MHNCUPANF 320) C adult male from Cuanavale River source lake, Moxico Province (
The following call description is based on a recording of a paratype male (
Based on our phylogenetic analysis, this species is currently confirmed from three widely separated localities and elevations ranging from near sea level to > 1400 m: i) Kissama National Park, on the outskirts of Angola’s capital city, Luanda, in coastal western Angola (Luanda Province); ii) central Angola (Bié Province); and iii) the source of the Cuanavale, Cuito, Cuando and Quembo rivers (Moxico Province). The identity of other known Angolan localities for Breviceps (black diamonds) remain uncertain without additional sampling and genetic data (Fig.
Breviceps ombelanonga differs from other species within the B. mossambicus group by net uncorrected mitochondrial p-distances of at least 9% (12S) and 11% (16S; Table
The preferred habitat for B. ombelanonga ranges from typical western Angolan savannah, with sandy soils and vegetation dominated by Adansonia digitata, Euphorbia conspicua, Acacia welwitschii and Combretum sp., together with a good grass coverage (
Photos of typical habitat of B. ombelanonga sp. nov.: A a view of the Kwanza River and bordering savannah, near the type locality, in Kissama National Park, Luanda Province B savannah near Embala Seque (14 km N of Cassumbi village), Bié Province C Cuanavale River source lake and associated miombo savannah woodland. Photographs by LMPC (A, B) and WC (C).
The name ombelanonga is a derived combination of two words in Umbundu, a native Angolan language, for rain (ombela) and frog (anonga). The species epithet is used as an invariable noun in apposition to the generic name.
Given that it appears widely distributed, we suggest that B. ombelanonga be included in the IUCN category of Least Concern. The type locality lies within Kissama National Park, which grants some legal protection from major habitat degradation and loss, though the park has recently experienced significant wildfires. Additionally, the paratype localities in southeastern Angola (visited during field activities related to the
Breviceps ombelanonga sp. nov. represents a phylogenetically distinct evolutionary lineage that is an Angolan endemic apparently geographically isolated from its closest congeners (Fig.
Further work is required to confirm the distributional range of B. ombelanonga, as well as whether it overlaps in distribution with either its sister taxon, B. poweri, or the more distantly related B. adspersus. Both occur in neighboring countries, B. poweri to the east/northeast (Zambia, Democratic Republic of Congo) and the B. adspersus to the south/southeast (Namibia, Botswana), and both have been suggested to occur in Angola (
We are not the first to recognize the lack of morphological variation within members of this anuran clade, which has led to historical taxonomic confusion and invoking hybridization for specimens that failed to conform to often scant descriptions of the type specimens (
As mentioned above, there is considerable genetic structure within B. ombelanonga, as well as among the four most closely related members of the B. mossambicus group (Fig.
We are grateful to Mariana P. Marques, Adam Ferguson, Ben Marks, John Cavagnaro, Philip Pastor, Suzana Bandeira, Ilola Jorge, Alvaro “Varito” Baptista, Ninda Baptista, Kerllen Costa, James Harvey, Roger Bills, Götz Neef, and Luke Verburgt for their invaluable field assistance, support, advice, and companionship. John Cavagnaro provided the life photo of the holotype. Mohamad Beidoun assisted with molecular work. This work was supported in part by the US National Science Foundation (DEB-1556255, 1556559 and DEB-1556585 to DCB, AMB, and MPH), the JRS Biodiversity Foundation (to DCB and AMB), and the National Geographic Society (Okavango Wilderness Project EC0715-15 to WC). The funders had no role in study design, data collection, and analysis, the decision to publish, or manuscript preparation. Field components of this work were facilitated by a Memorandum of Understanding between the University of Florida and The National Institute for Biodiversity and Protected Areas (INBAC) in Angola. Material used in this study was exported under the following permit numbers: 083/INBAC.MINAMB/2016 (to Villanova University) and 31/GGPCC/2016 (to the
Additional Breviceps material examined
Breviceps sp.: Angola: Lunda Sul Province: Alto Chicapa (
B. adspersus
: Botswana: Serowe (
B. mossambicus
: Malawi: Mount Mulanje (
B. poweri
: Democratic Republic of the Congo: Lualaba Province: Kalakundi (
Table S1. Morphological data used to perform PCAs
Data type: morphological data
Explanation note: Morphological data used to perform PCAs. See Table